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The two microaerophilic, Fe-oxidizing bacteria (FeOB)  Sideroxydans  ES-1 and  Gallionella  ES-2 have single circular chromosomes of 3.00 and 3.16 Mb that encode 3049 and 3006 genes, respectively. Multi-locus sequence analysis (MLSA) confirmed the relationship of these two organisms to one another, and indicated they may form a novel order, the Gallionellalaes, within the Betaproteobacteria. Both are adapted for chemolithoautotropy, including pathways for CO 2 -fixation, and electron transport pathways adapted for growth at low O 2 -levels, an important adaptation for growing on Fe(II). Both genomes contain Mto-genes implicated in iron-oxidation, as well as other genes that could be involved in Fe-oxidation. Nearly 10% of their genomes are devoted to environmental sensing, signal transduction, and chemotaxis, consistent with their requirement for growing in narrow redox gradients of Fe(II) and O 2 . There are important differences as well.  Sideroxydans  ES-1 is more metabolically flexible, and can utilize reduced S-compounds, including thiosulfate, for lithotrophic growth. It has a suite of genes for nitrogen fixation.  Gallionella  ES-2 contains additional gene clusters for exopolysaccharide production, and has more capacity to resist heavy metals. Both strains contain genes for hemerythrins and globins, but ES-1 has an especially high numbers of these genes that may be involved in oxygen homeostasis, or storage. The two strains share homology with the marine FeOB  Mariprofundus ferrooxydans  PV-1 in CO 2  fixation genes, and respiratory genes. In addition, ES-1 shares a suite of 20 potentially redox active genes with PV-1, as well as a large prophage. Combined these genetic, morphological, and physiological differences indicate that these are two novel species,  Sideroxydans lithotrophicus  ES-1 T  (ATCC 700298 T ; JCM 14762; DSMZ 22444; NCMA B100), and  Gallionella capsiferriformans  ES-2 T  (ATCC 700299 T ; JCM 14763; DSMZ 22445; NCMA B101).

Comparative genomics of freshwater Fe-oxidizing bacteria: implications for physiology, ecology, and systematics